Available online at www.sciencedirect.com
ScienceDirect
On the interaction of social affect and cognition:
empathy, compassion and theory of mind
1 1,2 1
Katrin Preckel , Philipp Kanske and Tania Singer
Empathy, compassion and Theory of Mind (ToM) are central which may be detrimental to the observer and to others
topics in social psychology and neuroscience. While empathy and compassion, on the other hand, which is a feeling of
enables the sharing of others’ emotions and may result in warmth and concern for the other. We conclude with a
empathic distress, a maladaptive form of empathic resonance, concise summary and an opinion statement.
or compassion, a feeling of warmth and concern for others,
ToM provides cognitive understanding of someone else’s
Defining and neurally characterizing empathy,
thoughts or intentions. These socio-affective and socio-
compassion and Theory of Mind
cognitive routes to understanding others are subserved by
Empathy describes the process of sharing feelings, that is,
separable, independent brain networks. Nonetheless they are
resonating with someone else’s feelings, regardless of
jointly required in many complex social situations. A process
valence (positive/negative), but with the explicit knowl-
that is critical for both, empathy and ToM, is self-other
edge that the other person is the origin of this emotion [1].
distinction, which is implemented in different temporoparietal
This socio-affective process results from neural network
brain regions. Thus, adaptive social behavior is a result of
activations that resemble those activations observed
dynamic interplay of socio-affective and socio-cognitive
when the same emotion is experienced first-hand (shared
processes.
network hypothesis) [2–5]. The first studies in neurosci-
ence targeting empathy investigated empathy in the
Addresses
1 domain of pain, showing that directly experiencing pain
Max Planck Institute for Human Cognitive and Brain Sciences,
Stephanstraße 1A, 04103 Leipzig, Germany and witnessing another person receiving painful stimuli
2
Institute of Clinical Psychology and Psychotherapy, Department of results in shared neural activations in the anterior insula
Psychology, Technische Universita¨ t Dresden, Dresden, Germany
(AI) and anterior middle cingulate cortex (aMCC) [6,7].
Meta-analyses have later identified these regions as a core
Corresponding author: Preckel, Katrin ([email protected])
network that is activated whenever we witness the suf-
fering of others [8,9]. Furthermore, this network is mod-
Current Opinion in Behavioral Sciences 2018, 19:1–6 ulated by individual differences in experienced negative
This review comes from a themed issue on Emotion-memory affect and empathy [6,10]. While empathy refers to an
interactions isomorphic representation of someone else’s affective
Edited by Mara Mather and Michael Fanselow state, compassion is a complementary social emotion
elicited by witnessing the suffering of others and is rather
For a complete overview see the Issue and the Editorial
associated with feelings of concern and warmth, linked to
Available online 31st July 2017
the motivation to help [2,11]. Empathy and compassion
http://dx.doi.org/10.1016/j.cobeha.2017.07.010
also differ on a neural level: compassion activates net-
ã
2352-1546/ 2017 The Authors. Published by Elsevier Ltd. This is an works that have previously been associated with reward
open access article under the CC BY-NC-ND license (http://creative-
and affiliation processes including the ventral striatum
commons.org/licenses/by-nc-nd/4.0/).
(VS), the nucleus accumbens, the ventral tegmental area
(VTA), the medial orbitofrontal cortex (mOFC) and the
subgenual anterior cingulate (sgACC)
As a social species, humans are continuously required to [12,13,14 ,15,16 ,17]. Congruently with these activations
process complex social signals. Successful multifaceted in reward-associated and affiliation-associated networks,
social interactions are enabled by socio-affective and compassion generates positive affect towards others’ suf-
socio-cognitive capacities such as empathy, compassion fering. In contrast to compassion, empathic distress,
and Theory of Mind (ToM). In this review we will first which is an alternative outcome of empathy, may be
define these social functions and describe the neural detrimental to the experiencer as well as to the suffering
networks associated with each of them. Further, we other [15,18].
discuss the interaction of empathy and ToM and delin-
eate the importance of one crucial process, that is, self- In contrast to socio-affective processes, socio-cognition
other distinction. Socio-affective and socio-cognitive pro- refers to taking another person’s perspective (also
cesses are also essential for how (prosocially) we interact referred to as ToM, mentalizing, or cognitive empathy).
with others, particularly when faced with others’ suffer- Rather than an emotional state, ToM yields abstract,
ing. Two possible outcomes of empathic sharing of propositional knowledge about the other’s mental state.
others’ suffering are empathic distress on the one hand, It describes the process of inferring and reasoning about
www.sciencedirect.com Current Opinion in Behavioral Sciences 2018, 19:1–6
2 Emotion-memory interactions
the beliefs, thoughts or emotions of others [19–22]. Cru- necessarily proficient mentalizers [27 ]. In line with such
cial brain regions involved in ToM include the ventral independent functions, selective impairments in empa-
temporoparietal junction (TPJ), superior temporal sulcus thy or ToM have been observed in different psycho-
(STS), temporal poles (TP), medial prefrontal cortex pathologies such as autism and psychopathy. In autism,
(MPFC) and precuneus/posterior cingulate (PCC) [23]. ToM is deficient [28–31], while no empathy deficits are
observed when controlling for alexithymia [32–35]. In
Interactions of social affect and social psychopathy or chronic aggression, in contrast, ToM is
cognition intact, but the propensity to empathize with others is
Both socio-affective and socio-cognitive processes have reduced [36,37].
been extensively investigated [6,8,9,23,24]. However,
research has only recently begun to study how these Despite being separable, empathy and ToM are jointly
processes are related and work together to achieve adap- required in many complex social situations. An indirect
tive social behavior. Making use of a novel task (Empa- source of evidence for such co-activation is a meta-analy-
ToM) [25 ] that stimulates both functions concurrently sis on different empathy for pain paradigms, which sug-
(see Figure 1), the respective neural correlates can be gests that the core region of the ToM network is co-
directly compared. The EmpaToM clearly activates sep- activated in empathy paradigms when additional inferring
arable brain networks that can be replicated in resting from a cue is required to understand the other’s state. [8].
state functional connectivity [25 ] and on a brain struc- The paradigms investigated in this meta-analysis varied
tural level [26 ]. Crucially, individual differences in depending on the information provided to the partici-
empathy and ToM are unrelated on a behavioral and pants. In picture-based paradigms, they were presented
neural level, that is, strong empathizers are not with visual depictions of someone in a painful situation,
Figure 1
EmpaToM task
Neutral Emotional Theory Factual nonToM ToM nonToM ToM of Mind Reasoning
Anna thinks that It is correct that
a) ... a) ...
b) ... b) ... c) ... c) ...
Valence Compassion Confidence
+ Anna + Question +
rating rating rating
1-3 s 2 s ~ 15 s 4 s 4 s 1-3 s max. 15 s 0-2 s 4 s How do you feel? How much compassion How confident do you
do you feel? feel?
negative neutral positive none very much uncertain certain
Current Opinion in Behavioral Sciences
The EmpaToM task. This figure presents the experimental set-up of the EmpaToM task (previously published here [25 ]). The design is a 2
(emotionality of video) Â 2 (ToM requirements) design, resulting in four different video types. Each actor presented each video type: emotional
negative and neutral videos, as well as videos with ToM requirements or factual reasoning demands. Each video was followed by various ratings,
a valence rating (ranging from negative to positive) and a compassion rating (ranging from none to very much compassion they felt for the person
in the video. Afterwards they answered a question that either probed for ToM or factual reasoning. After each question, participants were asked
how confident they felt about their answer. For further information on this task, please refer to Kanske et al. [25 ].
Current Opinion in Behavioral Sciences 2018, 19:1–6 www.sciencedirect.com
On the interaction of social affect and cognition Preckel, Kanske and Singer 3
whereas in cue-based paradigms, participants were pre- external world [39]. It has, however, also been discussed
sented a cue/hint that someone else is receiving painful as important in psychopathology, for instance in border-
stimulation. Common activations were observed in the line personality disorder for which a stress related men-
typical empathy related core network. However, the two talizing deficit has been hypothesized [40].
paradigm types resulted in different co-activations. As the
cue-based paradigms require additional inference of the Self-other distinction
other’s affective state from an abstract cue, the ToM The distinction of self and other constitutes an important
related network is co-active [8]. The pictorial presenta- element of both empathy and ToM, as it enables the
tion of someone in a painful situation, on the other hand, differentiation between one’s own emotional or mental
co-activates the action-observation network to decode the states and the states shared with others. Failure of self-
potential affective consequences of a certain action. other distinction results in a blending of these states,
These findings suggest that, dependent on the provided thereby inducing an egocentricity bias, the tendency to
information in a given context, different networks will be project one’s own emotional or mental state on someone
co-activated together with the core empathy-related else [41,42], or an altercentric bias, the influence of others’
network. states on judgments about oneself [43]. Cognitive ego-
centricity occurs when the own knowledge about a situa-
The two networks associated with empathy and ToM are tion influences the reasoning about what someone else
also jointly activated for complex evaluations of someone thinks about the situation [42], but can also affect simple
else’s feelings such as empathic accuracy, the ability to decisions about the visual perspective of another [44].
infer what someone else is feeling [38]. Here, participants Emotional egocentricity occurs when one’s own emo-
are asked to continuously rate another person’s emotions tional state influences the judgment of someone else’s
over an extended period of time; so the emotion of the affective state [41]. On a neural level, the right TPJ seems
other may be empathically shared, but it is also critical to to be critical to overcome cognitive egocentricity [45 ]. It
acquire abstract knowledge about that emotion to com- is also functionally connected to other parts of the ToM
plete the task accurately. neural network, such as the medial and ventrolateral PFC
and the PCC [45 ,46]. Overcoming emotional egocentric-
When activated together in complex social situations, the ity involves a slightly anterior temporoparietal region in
two networks may also directly influence each other. the right supramarginal gyrus (rSMG) [41]. Correspond-
Indeed, when confronted with others’ distress, the AI, ingly, disruption of rSMG activity using transcranial mag-
which is activated for negative affect sharing, inhibits netic stimulation leads to increased emotional egocen-
activity in TPJ, an important hub of the mentalizing tricity [41]. Interestingly, the SMG (see Figure 2) is also
network [27 ]. This reduction in TPJ activity is related part of a different functional connectivity network with
to impaired ToM performance [27 ]. The adaptive value extensive connections to the AI [47]. Moreover, increased
of this inhibitory mechanism in salient, emotional situa- coupling of SMG to dorsolateral PFC has been related to a
tions may be to prepare for immediate action in the reduction in emotional egocentricity in development
Figure 2
SMG aMCC PCC TPJ mPFC DLPF Theory of Mind GP Put STS VS AI NAcc Compassion sgACC mOFC TP VTA Empathy for Suffering SN
Current Opinion in Behavioral Sciences
Neural networks underlying empathy, compassion and Theory of Mind. The separable neural networks underlying empathy (blue), compassion
(red), and Theory of Mind (green) are presented. Abbreviations: AI, anterior insula; aMCC, anterior middle cingulate cortex; dlPFC, dorsolateral
prefrontal cortex; GP, globus pallidus; mOFC, medial orbitofrontal cortex; mPFC, medial prefrontal cortex; NAcc, nucleus accumbens; PCC,
posterior cingulate cortex; Put, putamen; sgACC, subgenual anterior cingulate cortex; SN, substantia nigra; STS, superior temporal sulcus; SMG,
supramarginal gyrus; TP, temporal poles; TPJ, temporoparietal junction; VS, ventral striatum; VTA, ventral tegmental area.
www.sciencedirect.com Current Opinion in Behavioral Sciences 2018, 19:1–6
4 Emotion-memory interactions
[46]. Thus, temporoparietal regions seem to play a crucial networks and compassion increases positive affect to
role in overcoming egocentricity, possibly contributing to buffer negative affect through the active generation of
self-other distinction. The activation of SMG and TPJ in positive emotions via reward-related and affiliation-
complex empathy and ToM paradigms, respectively related brain circuitries.
[25 ], corresponds with the crucial role of self-other
distinction for both capacities.
Concluding remarks
In the present paper, we have suggested that the social
Social behavior
brain is supported by different functions and underlying
Empathy and ToM provide us with an understanding of
neuronal networks and that it is important to differentiate
our interaction partners’ emotional and cognitive states,
between socio-affective versus socio-cognitive routes to
thereby forming the basis for flexible interactive behavior
the understanding of others. Sharing affective states with
and social decision making. Lately, neuroscientific
another person (empathy), feeling concern for another
research has focused on the relationship between social
(compassion) and reasoning about another person’s men-
emotions and social cognition in predicting prosocial
tal state (ToM) are separable on a conceptual, behavioral
behaviors [48 ,49]. For instance, using donation choices
and neural level; strong empathizers are not necessarily
for charity organizations, a recent study [48 ] showed
proficient mentalizers and each domain may be selec-
that an extensive cortical network is involved in forming
tively impaired in psychopathologies such as autism or
such prosocial decisions. Specifically, AI activity pre-
psychopathy. Furthermore, an original adaptive empathic
dicted generous donations when participants reported
response to the suffering of others can lead to a mal-
empathic feelings for the charity’s goal before, while
adaptive response, that is, empathic distress which can be
TPJ activity predicted donations, when participants took
reversed by learning how to turn empathy into compas-
the perspective of the organization’s cause [48 ].
sion. Compassion can thus be viewed as an emotion-
Remarkably, empathy and ToM measurements from
regulation strategy that buffers negative affect through
an independent task, were also related to these functions’
the active generation of positive affect relying on reward-
role in donation decisions, suggesting that stable individ-
related and affiliation-related brain circuitries. Despite
ual propensities are involved in prosocial behavior. In
the separability of socio-affective and socio-cognitive
contrast, impairments in empathy or ToM have been
functions, these social capacities are jointly activated
associated with antisocial behavior, for instance to hyper-
and interact in complex social situations. In synthesis,
aggression [18,37,50]. While empathy may be related to
the evidence supports a detailed view of the social mind,
prosociality, a particular outcome of empathy — empathic
not as a monolithic ‘social intelligence’, but as dynamic
distress — describes the intense sharing of other’s emo-
interplay between different functions and subserving
tions and may be detrimental, because it can result in own
neuronal networks that enable in different ways to engage
suffering as well as in asocial or even antisocial effects,
in prosocial behavior.
such as aggressive verbal behavior [18]. Short-term con-
templative training in compassion increases helping, even
when it is costly [51]. In contrast to empathic distress, Conflict of interest statement
compassion reduces punishment behavior in long-term Nothing declared.
compassion meditation practitioners even when they
were themselves the victim of fairness violations [52].
References and recommended reading
The critical property of compassion may be that it coun-
Papers of particular interest, published within the period of review,
teracts negative emotion elicited by experiencing others’ have been highlighted as:
suffering through positive emotion generation, thereby
of special interest
acting as an emotion regulation strategy [14 ,15,53]. The of outstanding interest
direct comparison of compassion to the most widely
1. de Vignemont F, Singer T: The empathic brain: how, when and
investigated emotion regulation strategy — reappraisal
why? Trends Cogn Sci 2006, 10:435-441.
— showed that in well-trained long-term meditators,
2. Singer T, Lamm C: The social neuroscience of empathy. Ann N Y
cognitive reappraisal was most efficient in reducing neg-
Acad Sci 2009, 1156:81-96.
ative affect, but compassion induction worked through
3. Gallese V: The roots of empathy: the shared manifold
up-regulation of positive affect and associated networks
hypothesis and the neural basis of intersubjectivity.
in the brain [14 ]. On a neural level, compassion was Psychopathology 2003, 36:171-180.
accompanied by activations in reward and positive emo- 4. Wicker B, Keysers C, Plailly J, Royet JP, Gallese V, Rizzolatti G:
Both of us disgusted in My insula: the common neural basis of
tion related networks [14 ]. Reappraisal on the other hand
seeing and feeling disgust. Neuron 2003, 40:655-664.
yielded activity in a fronto-parietal network associated
5. Decety J, Chaminade T: Neural correlates of feeling sympathy.
with cognitive control and attention regulation [54]. The
Neuropsychologia 2003, 41:127-138.
different brain activation patterns are, thus, in line with
6. Singer T, Seymour B, O’Doherty J, Kaube H, Dolan RJ, Frith CD:
the hypothesis that reappraisal down-regulates negative
Empathy for pain involves the affective but not sensory
affect mainly via executive and cognitive control components of pain. Science 2004, 303:1157-1162.
Current Opinion in Behavioral Sciences 2018, 19:1–6 www.sciencedirect.com
On the interaction of social affect and cognition Preckel, Kanske and Singer 5
7. Goubert L, Craig KD, Vervoort T, Morley S, Sullivan MJL, 25. Kanske P, Bo¨ ckler A, Trautwein FM, Singer T: Dissecting the
Williams ACD, Cano A, Crombez G: Facing others in pain: the social brain: introducing the EmpaToM to reveal distinct
effects of empathy. Pain 2005, 118:285-288. neural networks and brain-behavior relations for empathy and
theory of mind. Neuroimage 2015, 122:6-19.
8. Lamm C, Decety J, Singer T: Meta-analytic evidence for Adaptive behavior in complex social situations requires empathy, as well
common and distinct neural networks associated with directly as Theory of Mind (ToM). This study presents, for the first time, an
experienced pain and empathy for pain. Neuroimage 2011, experiment which tests both functions independently in the same indi-
54:2492-2502. vidual. Results reveal distinct networks for empathy and ToM and brain
activity in these networks predicts the corresponding behavior in these
9. Bzdok D, Schilbach L, Vogeley K, Schneider K, Laird AR,
two domains.
Langner R, Eickhoff SB: Parsing the neural correlates of moral
cognition: ALE meta-analysis on morality, theory of mind, and 26. Valk SL, Bernhardt BC, Bo¨ ckler A, Trautwein FM, Kanske P,
empathy. Brain Struct Funct 2012, 217:783-796. Singer T: Socio-cognitive phenotypes differentially modulate
large-scale structural covariance networks. Cereb Cortex
10. Birren JE, Casperson RC, Botwinick J: Pain measurement by the
2017, 27:1358-1368.
radiant heat method: individual differences in pain sensitivity,
This study investigated whether the separable social cognition networks
the effects of skin temperature, and stimulus duration. J Exp
underlying empathy and Theory of Mind is also distinguishable on a
Psychol 1951, 41:419-424.
structural level. Data from 270 healthy adults was included in the analysis
and the authors concluded that different routes of social cognition are
11. Goetz JL, Keltner D, Simon-Thomas E: Compassion: an
based on structural networks.
evolutionary analysis and empirical review. Psychol Bull 2010,
136:351-374.
27. Kanske P, Bo¨ ckler A, Trautwein FM, Parianen Lesemann FH,
Singer T: Are strong empathizers better mentalizers? Evidence
12. Klimecki OM, Leiberg S, Lamm C, Singer T: Functional neural
for independence and interaction between the routes of social
plasticity and associated changes in positive affect after
cognition. Soc Cogn Affect Neurosci 2016:1383-1392.
compassion training. Cereb Cortex 2013, 23:1552-1561.
By measuring both empathy and ToM in the same participants, this study
showed that participants’ capacities to empathize and mentalize are
13. Klimecki OM, Leiberg S, Ricard M, Singer T: Differential pattern
independent on a behavioral as well as on a neural level. Thus, strong
of functional brain plasticity after compassion and empathy
empathizers are not necessarily proficient mentalizers. However, in
training. Soc Cogn Affect Neurosci 2014, 9:873-879.
situations of high emotion, empathy-related neural activity can inhibit
ToM-related activation, leading to behavioral ToM impairments.
14. Engen HG, Singer T: Compassion-based emotion regulation
up-regulates experienced positive affect and associated
28. Frith U, Happe F: Autism spectrum disorder. Curr Biol 2005, 15:
neural networks. Soc Cogn Affect Neurosci 2015, 10:1291-1301.
R786-R790.
Proficient meditators and control participants watched short videos with
either negative or neutral emotional content during functional imaging.
29. Frith U: Cognitive explanations of autism. Acta Paediatr Suppl
Participants were asked to use compassion or reappraisal to regulate
1996, 416:63-68.
their emotions during video presentation. Both emotion regulation stra-
tegies decreased negative and increased positive affect. It is concluded
30. Baron-Cohen S, Leslie AM, Frith U: Does the autistic child have a
that compassion is an effective emotion regulation strategy, which, in
“theory of mind”? Cognition 1985, 21:37-46.
addition to downregulating negative affect (as is achieved in reappraisal),
mainly upregulates positive affect.
31. Happe FG: The role of age and verbal ability in the theory of
mind task performance of subjects with autism. Child Dev
15. Singer T, Klimecki OM: Empathy and compassion. Curr Biol
1995, 66:843-855.
2014, 24:R875-R878.
32. Bird G, Silani G, Brindley R, White S, Frith U, Singer T: Empathic
16. Weng HY, Fox AS, Hessenthaler HC, Stodola DE, Davidson RJ:
brain responses in insula are modulated by levels of
The role of compassion in altruistic helping and punishment
alexithymia but not autism. Brain 2010, 133:1515-1525.
behavior. PLOS ONE 2015, 10:e0143794.
The authors measured the social behavioral outcomes of compassion in
33. Bird G, Viding E: The self to other model of empathy: providing a
different contexts of economic games. Results show that compassion is
new framework for understanding empathy impairments in
more likely to lead to helping behavior towards a victim than to punish-
psychopathy, autism, and alexithymia. Neurosci Biobehav Rev
ment behavior towards a transgressor.
2014, 47:520-532.
17. Lutz A, Brefczynski-Lewis J, Johnstone T, Davidson RJ:
34. Gaigg SB, Cornell AS, Bird G: The psychophysiological
Regulation of the neural circuitry of emotion by compassion
mechanisms of alexithymia in autism spectrum disorder.
meditation: effects of meditative expertise. PLoS ONE 2008, 3:
Autism 2016.
e1897.
35. Hoffmann F, Koehne S, Steinbeis N, Dziobek I, Singer T:
18. Klimecki OM, Vuilleumier P, Sander D: The impact of emotions
Preserved self-other distinction during empathy in autism is
and empathy-related traits on punishment behavior:
linked to network integrity of right supramarginal gyrus. J
introduction and validation of the inequality game. PLOS ONE
Autism Dev Disord 2016, 46:637-648.
2016, 11:e0151028.
36. Meffert H, Gazzola V, den Boer JA, Bartels AA, Keysers C:
19. Premack D, Woodruff G: Does the chimpanzee have a theory of
Reduced spontaneous but relatively normal deliberate
mind. Behav Brain Sci 1978, 1:515-526.
vicarious representations in psychopathy. Brain 2013,
136:2550-2562.
20. Frith U, Frith CD: Development and neurophysiology of
mentalizing. Philos Trans R Soc B Biol Sci 2003, 358:459-473.
37. Winter K, Spengler S, Bermpohl F, Singer T, Kanske P: Social
cognition in aggressive offenders: impaired empathy, but
21. Saxe R, Kanwisher N: People thinking about thinking people —
intact theory of mind. Sci Rep 2017, 7.
the role of the temporo-parietal junction in “theory of mind”.
Neuroimage 2003, 19:1835-1842.
38. Zaki J, Weber J, Bolger N, Ochsner K: The neural bases of
empathic accuracy. Proc Natl Acad Sci U S A 2009, 106:11382-
22. Mitchell JP: The false dichotomy between simulation and
11387.
theory–theory: the argument’s error. Trends Cogn Sci 2005,
9:363-364 author reply 364.
39. Menon V, Uddin LQ: Saliency, switching, attention and control:
a network model of insula function. Brain Struct Funct 2010,
23. Schurz M, Radua J, Aichhorn M, Richlan F, Perner J:
214:655-667.
Fractionating theory of mind: a meta-analysis of functional
brain imaging studies. Neurosci Biobehav Rev 2014, 42:9-34.
40. Fonagy P, Luyten P: A developmental, mentalization-based
approach to the understanding and treatment of borderline
24. Singer T: The neuronal basis and ontogeny of empathy and
personality disorder. Dev Psychopathol 2009, 21:1355-
mind reading: review of literature and implications for future
1381.
research. Neurosci Biobehav Rev 2006, 30:855-863.
www.sciencedirect.com Current Opinion in Behavioral Sciences 2018, 19:1–6
6 Emotion-memory interactions
41. Silani G, Lamm C, Ruff CC, Singer T: Right supramarginal gyrus attention shifts differentially predict altruistic giving. J
is crucial to overcome emotional egocentricity bias in social Neurosci 2016, 36:4719-4732.
judgments. J Neurosci 2013, 33:15466-15476. Using a charitable donation task, the authors identified three separable
mechanisms (empathy, Theory of Mind (ToM), attention reorienting)
42. Royzman EB, Cassidy KW, Baron J: “I know, you know”:
subserving altruistic decision-making and linked them to separable neural
epistemic egocentrism in children and adults. Rev Gen Psychol
activations. Brain activity in the anterior insula and the temporoparietal
2003, 7:38-65.
junction corresponded to the respective socio-affective (empathy) or
socio-cognitive (ToM) process in which participants engaged when
43. Hoffmann F, Banzhaf C, Kanske P, Ga¨ rtner M, Bermpohl F,
making a decision. Donation choices predicted participants’ engagement
Singer T: Empathy in depression: egocentric and altercentric
in affective-processes or cognitive-processes.
biases and the role of alexithymia. J Affect Disord 2016, 199:23-
29.
49. Hare TA, Camerer CF, Knoepfle DT, Rangel A: Value
computations in ventral medial prefrontal cortex during
44. Samson D, Apperly IA, Braithwaite JJ, Andrews BJ, Bodley
charitable decision making incorporate input from regions
Scott SE: Seeing it their way: evidence for rapid and involuntary
involved in social cognition. J Neurosci 2010, 30:583-590.
computation of what other people see. J Exp Psychol Hum
Percept Perform 2010, 36:1255-1266.
50. Spenser KA, Betts LR, Das Gupta M: Deficits in theory of mind,
empathic understanding and moral reasoning: a comparison
45. Steinbeis N: The role of self-other distinction in understanding
between young offenders and non-offenders. Psychol Crime
others’ mental and emotional states: neurocognitive
Law 2015, 21:632-647.
mechanisms in children and adults. Philos Trans R Soc Lond B
Biol Sci 2016, 371:20150074.
51. Leiberg S, Klimecki OM, Singer T: Short-term compassion
This article comprehensively reviews the literature on self-other distinc-
training increases prosocial behavior in a newly developed
tion, cognitive and emotional egocentricity bias and the underlying neural
prosocial game. PLoS ONE 2011, 6:e17798.
systems, as well as their impact on social interactions.
52. McCall C, Steinbeis N, Ricard M, Singer T: Compassion
46. Steinbeis N, Bernhardt BC, Singer T: Age-related differences in
meditators show less anger, less punishment and more
function and structure of rSMG and reduced functional
compensation of victims in response to fairness violations.
connectivity with DLPFC explains heightened emotional
Front Behav Neurosci 2014, 8.
egocentricity bias in childhood. Soc Cogn Affect Neurosci 2015,
10:302-310.
53. Engen HG, Singer T: Affect and motivation are critical in
47. Mars RB, Sallet J, Schuffelgen U, Jbabdi S, Toni I, constructive meditation. Trends Cogn Sci 2016, 20:159-160.
Rushworth MFS: Connectivity-based subdivisions of the
54. Ochsner KN, Silvers JA, Buhle JT: Functional imaging studies of
human right “temporoparietal junction area”: evidence for
emotion regulation: a synthetic review and evolving model of
different areas participating in different cortical networks.
the cognitive control of emotion. Ann N Y Acad Sci 2012, 1251:
Cereb Cortex 2012, 22:1894-1903.
E1-E24.
48. Tusche A, Bo¨ ckler A, Kanske P, Trautwein FM, Singer T: Decoding
the charitable brain: empathy, perspective taking, and
Current Opinion in Behavioral Sciences 2018, 19:1–6 www.sciencedirect.com